In this series, we feature excerpts from biologist Sean Carroll’s book, Endless Forms Most Beautiful: The New Science of Evo Devo, a well-wrought account of how the field of evolutionary development biology, or Evo Devo, has shaped our understanding of how animals evolve, by studying their development. The Psalmist wrote of how God knits us together in our mother’s womb, and thanks to the methods and tools now available to biologists, these processes that were once hidden are being revealed through this exciting field. While we cannot attest to the author’s faith persuasion, Sean Carroll does an outstanding (and witty) job of making a complex set of ideas more manageable to understand, and we encourage you to read this book in its entirety.
In this post, Carroll introduces a bit of Evo Devo history, as well as the concept of how animal form can be understood as the product of two processes—development and evolution.
The editorial policy used in these excerpts can be found at the bottom of this post.
Embryos and Evolution
The first approach naturalists took to dealing with the great variety of animals was to sort them into groups, such as vertebrates (including fish, amphibians, reptiles, birds, and mammals) and arthropods (insects, crustaceans, arachnids, and more), but between and within these groups there are many differences. What makes a fish different from a salamander? Or an insect from a spider? On a finer scale, clearly a leopard is a cat, but what makes it different from a domestic tabby? And closer to home, what makes us different from our chimpanzee cousins?
The key to answering such questions is to realize that every animal form is the product of two processes—development from an egg and evolution from its ancestors. To understand the origins of the multitude of animal forms, we must understand these two processes and their intimate relationship to each other. Simply put, development is the process that transforms an egg into a growing embryo and eventually an adult form. The evolution of form occurs through changes in development.
Both processes are breathtaking. Consider that the development of an entire complex creature begins with a single cell—the fertilized egg. In a matter of just a day (a fly maggot), a few weeks (a mouse), or several months (ourselves), an egg grows into millions, billions, or, in the case of humans, perhaps 10 trillion cells formed into organs, tissues, and parts of the body. There are few, if any, phenomena in nature that inspire our wonder and awe as much as the transformation from egg to embryo to the complete animal. One of the great figures in all of biology, Darwin’s close ally Thomas H. Huxley, remarked:
The student of Nature wonders the more and is astonished the less, the more conversant he becomes with her operations; but of all the perennial miracles she offers to his inspection, perhaps the most worthy of admiration is the development of a plant or of an animal from its embryo.—Aphorisms and Reflections (1907)
The intimate connection between development and evolution has long been appreciated in biology. Both Darwin, inThe Origin of Species (1859) and The Descent of Man (1871), and Huxley in his short masterpiece, Evidence as to Man’s Place in Nature (1863), leaned heavily on the facts of embryology (as they were in the mid-nineteenth century) to connect man to the animal kingdom and for indisputable evidence of evolution. Darwin asked his reader to consider how slight changes, introduced at different points in the process and in different parts of the body, over the course of many thousands or a million generations, spanning perhaps tens of thousands to a few million years, can produce different forms that are adapted to different circumstances and that possess unique capabilities. That is evolution in a nutshell.
For Huxley, the nub of the argument was simple: we may marvel at the process of an egg becoming an adult, but we accept it as an everyday fact. It is merely then a lack of imagination to fail to grasp how changes in this process that are assimilated over long periods of time, far longer than the span of human experience, shape life’s diversity. Evolution is as natural as development. [SNIP]
While Darwin and Huxley were right about development as key to evolution, for more than one hundred years after their chief works, virtually no progress was made in understanding the mysteries of development. The puzzle of how a simple egg gives rise to a complete individual stood as one of the most elusive questions in all of biology. Many thought that development was hopelessly complex and would involve entirely different explanations for different types of animals. So frustrating was the enterprise that the study of embryology, heredity, and evolution, once intertwined at the core of biological thought a century ago, fractured into separate fields as each sought to define its own principles.
Because embryology was stalled for so long, it played no part in the so-called Modern Synthesis of evolutionary thought that emerged in the 1930s and 1940s. In the decades after Darwin, biologists struggled to understand the mechanisms of evolution. At the time of The Origin of Species, the mechanism for the inheritance of traits was not known. Gregor Mendel’s work was rediscovered decades later and genetics did not prosper until well into the 1900s. Different kinds of biologists were approaching evolution at dramatically different scales. Paleontology focused on the largest time scales, the fossil record, and the evolution of higher taxa. Systematists were concerned with the nature of species and the process of speciation. Geneticists generally studied variation in traits in just a few species. These disciplines were disconnected and sometimes hostile over which offered the most worthwhile insights into evolutionary biology. Harmony was gradually approached through an integration of evolutionary viewpoints at different levels. Julian Huxley’s book Evolution: The Modern Synthesis (1942) signaled this union and the general acceptance of two main ideas. First, that gradual evolution can be explained by small genetic changes that produce variation which is acted upon by natural selection. Second, that evolution at higher taxonomic levels and of greater magnitude can be explained by these same gradual evolutionary processes sustained over longer periods.
The Modern Synthesis established much of the foundation for how evolutionary biology has been discussed and taught for the past sixty years. However, despite the monikers of “Modern” and “Synthesis,” it was incomplete. At the time of its formulation and until recently, we could say that forms do change, and that natural selection is a force, but we could say nothing about how forms change, about the visible drama of evolution as depicted, for example, in the fossil record. The Synthesis treated embryology as a “black box” that somehow transformed genetic information into three-dimensional, functional animals.
The stalemate continued for several decades. Embryology was preoccupied with phenomena that could be studied by manipulating the eggs and embryos of a few species, and the evolutionary framework faded from embryology’s view. Evolutionary biology was studying genetic variation in populations, ignorant of the relationship between genes and form. Perhaps even worse, the perception of evolutionary biology in some circles was that it had become relegated to dusty museums.
Such was the setting in the 1970s when voices for the reunion of embryology and evolutionary biology made themselves heard. Most notable was that of Stephen Jay Gould, whose book Ontogeny and Phylogeny revived discussion of the ways in which the modification of development may influence evolution. Gould had also stirred up evolutionary biology when, with Niles Eldredge, he took a fresh look at the patterns of the fossil record and forwarded the idea of punctuated equilibria—that evolution was marked by long periods of stasis (equilibria) interrupted by brief intervals of rapid change (punctuation). Gould’s book and his many subsequent writings reexamined the “big picture” in evolutionary biology and underscored the major questions that remained unsolved. He planted seeds in more than a few impressionable young scientists, myself included.
To me, and others who had been weaned on the emerging successes of molecular biology in explaining how genes work, the situations in embryology and in evolutionary biology were both unsatisfying, but they presented enormous potential opportunities. Our lack of embryological knowledge seemed to turn much of the discussion in evolutionary biology about the evolution of form into futile exercises in speculation. How could we make progress on questions involving the evolution of form without a scientific understanding of how form is generated in the first place? Population genetics had succeeded in establishing the principle that evolution is due to changes in genes, but this was a principle without an example. No gene that affected the form and evolution of any animal had been characterized. New insights in evolution would require breakthroughs in embryology.